ABHANDLUNGEN DER GEOLOGISCHEN BUNDESANSTALT AbhGeolB A ,GA - - ISSN 0378-08641 ISBN 3-85316-007-7 Band 56/2 S. 669-678 I Wien, Dezember 1999 Geologie ohne Grenzen Redaktion: Festschrift 150 Jahre Geologische Bundesanstalt Harald Lobitzer & Pavol Grecula

Diachronous deposits of Lake Pannon in the Kisalföld Basin reflect basin and mollusc evolution

IMRE MAGYAR, PÄL MÜLLER, DANA H. GEARY, HILARY C. SANDERS and GABOR C. TARI

Pannonischer See Kleine Ungarische Tiefebene Slowakei Österreichische Karte 1:50 000 Beckenanalyse Blätter 60, 61, 62, 77, 78, 79, 80, 107, 108, 109, 137, 138, 139, 167, 168 Mollusken-Evolution

Contents

Zusammenfassung 669 Abstract 670 1. Geological setting 670 2. Materials and methods 671 3. Comparison of the Burgenland and TCR faunas 672 3.1. Similarities 672 3.2. Differences 672 4. Discussion 675 Acknowledgements 675 References 675 Appendix 676

Diachrone Ablagerungen des Pannonischen Sees in der Kleinen Ungarischen Tiefebene als Zeugen der Beckenentwicklung und der Molluskenevolution

Zusammenfassung

Das jungtertiäre Becken der Kleinen Ungarischen Tiefebene (Kisalföld) bildet einen Teilabschnitt des mitteleuropäischen Pannonischen Beckens und umfaßt Gebiete in der südwestlichen Slowakei, im östlichen Österreich sowie im nordwestlichen Ungarn. Littorale Sedimente des spätmiozänen Pannonischen Sees sind sowohl an dessen Westrand im Burgenland aufgeschlossen, als auch an dessen Ostrand im Transdanubischen Mittelgebirge. Reflexionsseismische Profile bezeugen, daß die Sedimentfüllung der Kleinen Ungarischen Tiefebene von NW nach SE progradierte. Als Folge dessen stellen die littoralen Sedimente des Burgenlands und des Transdanubischen Mittelgebirges nicht gegenüberliegende Küstenbildungen der Kleinen Ungarischen Tiefebene dar, sondern wurden entlang der gleichen progradierten nordwest­ lichen Küstenlinie des Pannonischen Sees, jedoch zu jeweils anderer Zeit, abgelagert. Die Molluskenfauna der westlichen Aufschlüsse ist einförmig ausgebildet und wird hier als "Fauna des Burgenlands" bezeichnet. Gleichermaßen ist auch die Fauna der östlichen Aufschlüsse recht einförmig ausgebildet und wird hier als "Fauna des Transdanubischen Mittelgebirges" bezeichnet. Aufgrund der Ähnlichkeiten zwischen diesen beiden Faunen, kann wohl davon ausgegangen werden, daß ihre Lebensräume im littoralen Bereich des Pannonischen Sees sehr ähnlich ausgebildet waren. Mehrere Bivalvenarten zeigen jedoch deutliche morphologische Unterschiede in den beiden Faunengebieten, die auf evolutive Ursachen zurückgeführt werden. Der Unterschied zwischen der Fauna des Burgenlands und der Fauna des Transdanubischen Mittelgebirges hat biostratigraphische Bedeutung, wodurch die Grenze zwischen der Lymnocardium conjungens Zone (Burgenland) und der Lymnocardium ponticum Zone (Transdanubisches Mittelgebirge) definiert wird. Magnetostratigraphische Daten legen nahe, daß mit einem Zeitunterschied im Bereich von etwa 500.000 Jahren zu rechnen sein dürfte.

Addresses of the authors: IMRE MAGYAR, MOL Hungarian Oil & Gas Company, Satthyäny u. 45, H-1039, Budapest; PÄL MÜLLER, Geological Institute of , Stefänia tit 14, H-1143, Budapest; DANA H. GEARY, HILARY C. SANDERS, Department of Geology & Geophysics, University of Wisconsin-Madison, 1215 Dayton Street, Madison, Wl 53706, U.S.A.; GABOR C. TARI, BP Amoco Exploration, 501 West Lake Park Boulevard, Houston, TX 77079-2696, U.S.A.

669 Abstract

The Neogene Kisalföld Basin is a subunit of the Central European Pannonian Basin. It includes southwestern Slovakia, the easternmost part of , and northwestern Hungary. Littoral sediments of the Late Miocene Lake Pannon outcrop along the western (Burgenland) and eastern (Transdanubian Central Range, TCR) margins of the basin. Seismic reflection profiles across the Kisalföld Basin indicate that the basin was filled with sediments via progradation from the NW to the SE. Consequently, the littoral sediments of Burgenland and the Transdanubian Central Range do not represent opposite shores of the Kisalföld Basin; instead, they were deposited on the same progra- ding northwestern shoreline of Lake Pannon during two different time intervals. The mollusc fauna of the western outcrops is uniform (refer­ red to here as the Burgenland fauna). Similarly, the eastern outcrops yield a uniform fauna (the TCR fauna). Similarity between the Burgenland and the TCR faunas allows us to establish that they lived in similar environments in the littoral zone of Lake Pannon. There are, however, consistent morphological differences between Burgenland and TCR forms in several bivalve species, which are thus interpreted as evolutionary change. The difference between the Burgenland and TCR faunas is useful for biostratigraphy; it defines the boundary be­ tween the Lymnocardium conjungens Zone (Burgenland) and Lymnocardium ponticum Zone (TCR). Magnetostratigraphic data suggest that the time difference involved is on the order of a half million years.

1. Geological setting the littoral zone of Lake Pannnon shifted from the NW to­ wards the SE (Text-Fig. 2). Seismic profiles also suggest that The Kisalföld ("Danube") Basin is a subbasin of the at this time the Transdanubian Central Range (TCR) was a Neogene Pannonian Basin of Central . Its area is submerged sill, or a series of islands at best. shared by Austria, Slovakia, and Hungary (Text-Fig.1). The Due to Pliocene-Quaternary tectonic inversion, the sedi­ geology of the Kisalföld basin has recently been treated by mentary formations of Lake Pannon became exposed and KÖRÖSSY (1987), KEITH et al. (1994), TARI (1994, 1996), eroded along the western and eastern edges of the basin, HORVÄTH et al. (1995a), MATTICK et al. (1996), HRUSECKY et al. while the central part underwent continued subsidence, and (1996), and KOVÄC & BARÄTH (1996). The Kisalföld Neogene was covered by Pliocene and Quaternary terrestrial sedi­ Basin is superimposed on the Alpine thrust-fold belts of the ments (HORVÄTH et al., 1995b). Along the western edge of Eastern Alps and Western Carpathians. Following an Early to the basin, the littoral sediments of Lake Pannon outcrop in Middle Miocene, partly terrestrial, partly marine synrift stage, a number of fossiliferous localities in Burgenland and adja­ the bulk of the Neogene sedimentary formations were de­ cent areas of Slovakia and Hungary, from Pezinok in the posited in the Late Miocene brackish Lake Pannon, which north to Stegersbach in the south. Similar sediments out­ flooded the area due to a widespread post-rift (thermal) sub­ crop at the eastern boundary of the basin, along the western sidence. Seismic reflectors show that the Kisalföld Basin was foot of the TCR, from Chl'aba in the north to Ukk in the south filled with sediments prevailingly from the NW, implying that (Text-Fig. 1).

Text-Fig. 1. The Kisalföld Basin with locations referred to in the text and in the Appendix.

670 NW SE

QUATERNARY 10 KM

PALEOZOIC (LOWER/MIDDLE AUSTROALPINE NAPPES)

3- MESOZOIC/PALEOZOIC (TRANSDANUBIAN CENTRAL RANGE)

4-A Text-Fig. 2. Line drawing interpretation of a seismic profile illustrating progradation from the northwest towards the south­ east across the Kisalföld Basin. For location see Fig. 1. Vertical exaggeration is ten-fold at 2000 m/s velocity. For the original seismic profile see TARI (1994) and HORVÄTH et al. (1995b).

We observed that the littoral bivalve fauna from the western JPL: Collection of JOSEF PAUL LUEGER (St. Leonhard am Forst, margin of the basin ("Burgenland fauna") is uniform in the Austria) sense that it represents a particular evolutionary stage in each MÄFI: Collection of the Hungarian Geological Institute (Budapest, lineage. We found that the same is true for the eastern margin Hungary) as well; apart from slight facies differences, any member of MM: Collection of PAL MÜLLER and IMRE MAGYAR (Budapest, this "TCR fauna" is represented by the same evolutionary stage Hungary) in the outcrops throughout the entire length of the TCR. NHMW: Collection of the Vienna Natural History Museum (Vienna, There are striking similarities between these uniform western Austria) and eastern faunas, suggesting that their paleoenvironments TTM: Collection of the Hungarian Natural History Museum were very similar. In a number of bivalve species, however, (Budapest, Hungary) UV: Collection of the University of Vienna (Vienna, Austria) there are relatively slight, but consistent morphological differ­ ences between the two faunas, reflecting evolutionary chan­ ges through time. These differences in most cases are easily The fossils were collected from littoral sandy or gravely de­ recognizable, and make the two faunas distinguishable. posits. Even in the case of museum specimens, we were able to identify the original embedding rock because we knew the outcrop from which the specimens came, the description 2. Materials and methods of the outcrop was available, or because sand grains attached to the shells indicated the sandy environment. These sand lay­ In this study, we included only fossils that we have seen ers are either amalgamated, forming thick sandy sequences, ourselves or that were depicted in publications. We did not or they are thin intercalations in sublittoral silt sequences. use reports or determinations that we have not had the op­ In Burgenland as well as adjacent areas of Slovakia and portunity to check. We utilized the following museum and pri­ Hungary, the littoral deposits of Lake Pannon are of various vate collections (the list of fossils that formed the basis for ages. In this study we included the youngest and most wide­ this study is given in the Appendix): spread littoral deposits only, omitting older formations (Congeria ornithopsis and Congeria hoernesiZones, or B and C BL: Collection of the Burgenland Museum (Eisenstadt, Austria) Zones of PAPP, 1951) which outcrop in more restricted areas (e. DHG: Collection of DANA H. GEARY (Madison, USA) (Stegersbach g. in several localities around Sopran [PAPP, 1951, VITÄLIS, 1951], material is courtesy of FRANZ SAUERZOPF) and in the Styrian Basin [EBNER & SACHSENHOFER, 1995]). These FS: Collection of Franz Sauerzopf (Rust, Austria) older formations are missing at the eastern margin of trie basin. HPM: Collection of the Croatian Natural History Museum (Zagreb, Our objective here is not to compile a comprehensive list Croatia) of bivalve species for each locality; instead, we want to com-

671 plete earlier published information with unpublished data co­ differences in the frequency of certain species exist as well. ming from different collections, in order to demonstrate simi­ "Didacna" deserta is very common in Burgenland, but we larities and differences among bivalve faunas. We omitted know only a single intact specimen from the TCR. Congeria some relatively rare, or less studied dreissenid and cardiid ungulacaprae, on the other hand, is one of the most common species, and the sporadically appearing Pisidium. On the other species in the TCR, but it is much less common in Burgenland. hand, we included species that rarely occur in the given fau­ Aside from these conspicuous taxonomic differences, na if they are common on the opposite side of the basin (e. g. slight but consistent morphological differences characterize Congeria ungulacaprae is common in the TCR fauna, so we the ancestor (Burgenland) - descendant (TCR) species included its occurrences in Sopron and Pezinok; or pairs in some bivalves. Traditionally, the ancestral and the de­ "Didacna" deserta is very common in the Burgenland fauna, rived forms were described as different species, although the so we included its single known occurrence in the TCR area). morphological difference between them is sometimes very slight, and can be appreciated only when larger samples are compared. Comparison of the Burgenland In Burgenland specimens of Unio atavus, the position of and TCR faunas the beak in relation to the anterior margin is variable, but ty­ pically quite distant. In its descendant, U. mihanovici, however, it is invariably terminal, very close to the anterior margin. 3.1 Similarities In addition, the beak is always above the dorsal margin in U. atavus, whereas it is in much lower position in U. miha­ Many identical or very closely related species occur in the novici (MÜLLER, 1990; Plate 1, Fig. A, A'). Burgenland and TCR faunas. Some of these common species According to PAPP (1950), the Vienna Basin and Burgenland are rare in other parts of the Pannonian basin, thus enhancing fossil record shows that Congeria lost its apophyse (a tiny the impression that the Burgenland and TCR faunas are plate where the adductor muscle attached) in at least two inde­ closely related. Some of our new findings confirm a close re­ pendent lineages. Congeria has a well-developed apophyse, lationship between the two faunas: we found Euxinicardium whereas the lack of it is a diagnostic feature of Dreissena, schreien, a species that had been known from the TCR only, where both the anterior adductor and retractor attach to the in Burgenland. We also found two "typical" Burgenland spe­ same septum. One of these morphological changes is reflec­ cies, Lymnocard'tum tucani and "Monodacna" viennensis, in ted in the Burgenland/TCR material. Congeria gitneri (= "D. the TCR. minima" in LUEGER, 1980) in Stegersbach and Großhöflein Species that occur in both faunas include: Congeria ungu­ (Burgenland) has a rudimentary apophyse (Plate 1, Fig. B, C, lacaprae, Dreissena bipartita, Lymnocardium hantkeni, L. tu­ D). In Dreissena auricularis (TCR), however, there is no trace cani, Caladacna steindachneri, Euxinicardium schreteri, of the apophyse left (Plate 1, Figs. B', C, D'). (We know a "Monodacna" viennensis, "Didacna" deserta, and Parvi- sample from Pezinok [MÄFI PI. 6091] where there is no apo­ dacna sp. Ancestor - descendant pairs in the two faunas are physe, although this locality yielded Burgenland fauna.) (with the older, Burgenland form first): Unio atavus-U. miha- The Burgenland Lymnocardium schedelianum is a large novici, Congeria gitneri-Dreissena auricularis, Lymno­ cockle with relatively wide and rounded ribs, which are always cardium schedelianum-L. variocostatum, L. conjungens-L. separated by intercostal space (Plate 1, Fig. E). The only penslii, and L. edlaueri-L. ponticum. difference between this form and its descendant from the TCR, L. variocostatum, is that the ribs of the latter are com­ pletely flat and tightly spaced; intercostal space is present 3.2. Differences only on the very anterior slope of the shell (Plate 1, Fig. E'). Lymnocardium conjungens (Burgenland; Plate 1, Fig. F, Some of the differences between the Burgenland fauna and G) is similar to the early form of L penslii (TCR). The latter the TCR fauna are conspicuous. The TCR fauna lacks shoul­ is generally larger, and with different proportions in being dered Melanopsis (M. vindobonensis, M. fossilis), certain higher, and having a wider posterior gape (Plate 1, Fig. F', large Congeria species (C. pancici, C. spathulata), and some G'). The umbo above the hinge is also higher. LUEGER (1980) cardiid species (L. brunnense, L danicici, L. carnuntinum, L reported the common occurrence of L. conjungens and L. stoosi), whereas the Burgenland fauna lacks Dreissenomya pensliiimm Großhöflein, Burgenland. Indeed, his "L. penslii" unioides, "Lymnocardium"priscae, and L. apertum. Significant specimens are unusually large for L. conjungens, but they

Plate 1

Ancestor - descendant pairs in bivalves from the western (Burgenland) and eastern (TCR) margins of the Kisalföld Basin. Fig. -A: Unio atavus Hörnes, Großhöflein. Fig. -A': Unio mihanovici BRUSINA, Däka; Fig. -B, C, D: Congeria gitneri BRUSINA, B, C: Stegersbach, D: Großhöflein. Fig. -B', C, D' Dreissena auricularis (FUCHS), B': Läzi, C': Szäk, D': Neszmely. Fig. -E: Lymnocardium schedelianum (PARTSCH) (authorship and taxonomic status needs revision), Oggau. Fig. -E': Lymnocardium variocostatum VITÄLIS, Däka. Fig. -F, G: Lymnocardium conjungens (HÖRNES), Stegersbach. Fig. -P, G': Lymnocardium penslii (FUCHS), F': Ukk, G': Tüskevär. Fig. -H, I, J: Lymnocardium edlaueri PAPP, H: Pezinok, I, J: Stegersbach. Fig.-H', I', J': Lymnocardium ponticum HALAVÄTS, H': Papa, I': Nyäräd, J': Däka.

Scale bars indicate 1 cm, except Fig.-B, C, B', D', where it is 1 mm, and Fig.-D and C, where it is 100 \im.

672 673 Text-Fig. 3. Generalized cartoon representing the paleogeographic evolution of the Kisalföld Basin between the time of the "Burgenland fauna" (Lymnocardium conjungens Biochron; Fig. A) and the time of the "TCR fauna" {Lymnocardium ponticum Biochron; Fig. B).

Text-Fig. 4. Assessment of the age of the Burgenland and TCR faunas by means of magnetostratigraphy. Along the western margin of the Kisalföld Basin, the boundary between fine-grained, sublittoral slope deposits and delta front sands were correlated to the top of C5n (Nagylozs-1, Zsira-1) and to the base of C4Ar2r (Szombathely-ll). These correspond to ca. 9.7 and 9.5 Ma, respectively, according to BERGGREN et al. (1995). The littoral lacustrine fauna appears both in the Nagylozs-1 and Szombathely-ll boreholes in C4Ar2r. At the eastern margin of the basin, in the Duka-ll borehole, the base of the delta front deposits correlates to C4Ar1n (ca. 9.3 Ma), and the littoral fauna occurs from C4Ar1n to C4n. (Correlation of the actual polarity record of each borehole to the standard scale was carried out by Lantos, using the long normal C5n as tie-point in the western boreholes, and utilizing seismic profiles to connect the Duka section with Szombathely; see also MAGYAR et al., 1999.)

674 are injured and fragmentary thus cannot be subject to mor­ evolved. The Burgenland fauna, however, could live in the TCR phometry analysis. region, in littoral zones around islands. Indeed, we know one Lymnocardium edlaueri is a common, but small bivalve in such occurrence from the southernmost part of the TCR. The Burgenland (Plate 1, Fig. H, I, J) that is very similar to Kalla Gravel and Sand Formation (JÄMBOR, 1980) contains ty­ L. ponticum (TCR; Plate 1, Fig. H', I', J1). L edlaueri tends to pical "Burgenland" molluscs: Unio atavus, Congeria sp. ex be more elongate antero-posteriorly; L ponticum usually group C. subglobosa-pancici, Lymnocardium cf. soproniense (or grows bigger, and in larger specimens the umbo is high schedelianum?), and Melanopsis fossilis (see MAGYAR, 1988). above the hinge. There are no radioisotopic ages from the Kisalföld Basin to In two additional cases, the significance of the morpholo­ determine the age of the respective faunas. We must rely on gical differences between the Burgenland and the TCR spe­ magnetostratigraphic analyses from the Nagylozs-1, Zsira-1, cimens is unclear because we have so few specimens from Szombathely-ll, and Duka-ll boreholes. Interpretation of these the TCR. Lymnocardium tucanifrom Burgenland usually has polarity profiles by LANTOS (in LANTOS & ELSTON, 1995 and pers. more than 30 ribs. The TCR specimens, however, have only comm.) is anchored to the long-lasting Chron 5 normal. In the 23 to 26. Apart from this difference, the shells from the two Nagylözs and Szombathely boreholes, which lie close to the regions look very much alike. The only known full TCR spe­ western margin of the Kisalföld Basin, the interval with the litto­ cimen of "Didacna" deserta significantly differs from the ral lacustrine fauna was correlated to C4Ar2r (MAGYAR et al., Burgenland specimens in having a longer posterior margin, 1999; cf. KORPÄS-HÖDI, 1992 and LANTOS & ELSTON, 1995). On and consequently, a smaller angle between the posterior the opposite (eastern) side of the basin, in the Duka-ll bore­ and posterodorsal margins. The ventralmost point of the hole, however, the littoral fauna correlates to C4Ar1 r (MAGYAR valve is in the posterior part, instead of being in central et al., 1999; identification of molluscs in all three boreholes position (see MILAN et al., 1974). was carried out by KORPÄS-HÖDI; Text-Fig. 4). If we accept this correlation, then the age of the Burgenland fauna is somewhat older than 9.5 mys (because the Burgenland outcrops lie west­ 4. Discussion ward from these boreholes), whereas the age of the TCR fau­ na is slightly younger than 9.3 mys. The age difference be­ The only alternative to an evolutionary explanation for the tween the western and eastern littoral faunas of the Kisalföld morphological differences observed between Burgenland Basin is thus on the order of 0.5 million years. and TCR bivalves is that these differences reflect geographic variation. This hypothesis would require the presence of a paleogeographic barrier between the western and eastern Acknowledgements margins of the Kisalföld Basin, such as, for instance, a pro- foundal lacustrine zone in the axis of the basin. Seismic evi­ We thank JOSEF PAUL LUEGER and FRANZ SAUERZOPF for making it dence, however, indicates that although the Burgenland possible for us to study the fossils they had collected in Burgenland. outcrops and the TCR outcrops are presently situated on the We acknowledge the kind help of ORTWIN SCHULTZ (Natural History opposite margins of the Kisalföld Basin, historically they re­ Museum, Vienna), FRITZ STEININGER (formerly University of Vienna), present the same (NW) shoreline of Lake Pannon at two dif­ MARIA TSCHACH (Eisenstadt), JÄNOS SZABÖ, ALFRED DULAI (Hungarian ferent times in the evolution of the lake (Text-Fig. 3). Natural History Museum, Budapest), LÄSZLÖ KORDOS, ENDRE KROLOPP (Geological Institute of Hungary, Budapest), ZLATA JURISIC- The morphological and temporal differences between the POLSAK and KATARINA KRIZMANIC (Croatian Natural History Museum, Burgenland fauna and the TCR fauna provide the basis for Zagreb) for assistance with museum collections. This publication the distinction of the Lymnocardium conjungens Zone was sponsored by the National Science Foundation (EAR- (Burgenland) and L. ponticum Zone (TCR). Our paleogeo­ 9706230), by the U. S. - Hungarian Science & Technology Joint graphic model implies that between the Burgenland and Fund under project JFNo 511, by the Hungarian CTKA (project # T TCR faunas there must be a continuous littoral fossil record 019679), and by the Albert and Alice Weeks Fund of the Department buried beneath Pliocene and Quaternary sediments in the of Geology & Geophysics, University of Wisconsin-Madison. central part of the Kisalföld Basin. Our progradational model has further predictions/consequ­ ences. It implies that 1) the youngest lacustrine fauna in the References Vienna Basin is older than the Burgenland fauna; and 2) the littoral fauna southeast of the TCR is younger than the TCR ANDRUSOV, N.: Fossile und lebende Dreissensidae Eurasiens. - fauna. The second prediction is met by the presence of the Travaux de la Societe des Naturalistes de St.-Petersbourg. Section Lymnocardium decorum Zone (with Tihany- and de Geologie et de Mineralogie, 25, 1-683, avec un atlas de vingt planches, Sankt Petersburg 1897. Rädmänesti-type faunas; see MÜLLER, 1990 and MARINESCU, BARTHA, F.: Läzi felsö-pannöniai korüfaunäjänak biosztratigräfiai vizsgälata 1990) south and east of the TCR. Data relevant to the first (Depouillement biostratigraphique de la faune pannonienne supe- prediction are more ambiguous. Austrian, Czech, and Slovak rieure de la localite Läzi). - Annual Report of the Hungarian stratigraphers traditionally assign the uppermost lacustrine Geological Institute of 1960, 265-283, Budapest 1963. fauna of the Vienna Basin as well as that of Burgenland to BERGGREN, W.A., KENT, D.V., SWISHER, C.C., & AUBRY, M-P.: A revised Zones D and E of PAPP (1951), and do not recognize an age Cenozoic geochronology and chronostratigraphy. - In: BERGGREN, W.A., KENT, D.V., AUBRY, M-P. & HARDENBOL, J. (Eds): Geochrono­ difference between the faunas of the Vienna Basin and logy time scales and global stratigraphic correlation, SEPM Special Burgenland (see, for instance, PAPP, 1951; SAUERZOPF, 1952; Publication, 54, 129-212, 1995. LUEGER, 1980; JiRiCEK, 1985; FORDINÄL, 1997). At this time, BRUSINA, S.: Iconographia molluscorum fossilium in tellure Tertiana we cannot provide convincing paleontological evidence that Hungariae, Croatiae, Slavoniae, Dalmatiae, Bosniae, Herzegovinae, the Vienna Basin littoral faunas are older than those from Serbiae et Bulgariae inventorum. Atlas II, Plates 1-30, Zagreb 1902. Burgenland, however we know of no evidence that argues EBNER, F. & SACHSENHOFER, R.F.: Paleogeography, subsidence and against this interpretation. thermal history of the Neogene Styrian Basin (Pannonian Basin sys­ tem, Austria). - Tectonophysics, 242, 133-150, Amsterdam 1995. According to this progradational model, the TCR lacustrine FORDINÄL, K: Congeria doderleini Brusina from Pannonian sediments fauna cannot occur in Burgenland, because the latter area was in a loam pit of the brick plant at Pezinok. - Zapad. Karpaty, Ser. transformed into alluvial plains by the time the TCR fauna Paleont, 15, 71-77, Bratislava 1991.

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(Feherpart) in Hungary. - In: STEVANOVIC, P.M., NEVESSKAJA, L.A., FUCHS, Th.: Beiträge zur Kenntnis fossiler Binnenfaunen 4 und 5: Die MARINESCU, Fl., SOKAC, A. & JÄMBOR, Ä. (Eds.): Chronostratigraphie Fauna der Congerienschichten von Tihany am Plattensee und Küp und Neostratotypen. Neogen der Westlichen ("Zentrale") Paratethys bei Papa in Ungarn. - Jahrbuch k. k. Geol. Reichsanstalt, 20, VIII, PI1, Pontien, 558-581. JAZU and SANU, Zagreb-Beograd 1990. 531-548, Wien 1870. PAPP, A.: Übergangsformen von Congeria zu Dreissena aus dem HORVÄTH, F., TARI, G. & BOKOR, CS. (Eds.) (a): Hungary: Extensional Pannon des Wiener Beckens. - Annalen des Naturh. Museums collapse of the Alpine orogene and hydrocarbon prospects in the Wien, Wien 1950. basement and basin fill of the western Pannonian Basin. - AAPG PAPP, A.: Das Pannon des Wiener Beckens. - Mitteilungen der International Conference and Exhibition, Field Trip Notes 6, 1-204, Geologischen Gesellschaft in Wien, 39-41, 99-193, Wien 1951. Nice 1995. PAPP, A.: Die Molluskenfauna des Pannon im Wiener Becken. - HORVÄTH, F., CLOETINGH, S. & TARI, G. (b): Stress-induced late stage Mitteilungen der Geologischen Gesellschaft in Wien, 44, 85-222, subsidence anomalies of the Pannonian Basin. - In: HORVÄTH, F., Wien 1953. TARI, G. & BOKOR, CS. (Eds.): Hungary: Extensional collapse of the PAPP, A.: Gastropoda (Neritidae, Viviparidae, Valvatidae, Hydrobiidae, Alpine orogene and hydrocarbon prospects in the basement and Stenothyridae, Truncatellidae, Bulimidae, Micromelaniidae, Thiari- basin fill of the western Pannonian Basin. AAPG International dae) und Bivalvia (Dreissenidae, Limnocardiidae, Unionidae) des Conferenceand Exhibition, Field Trip Notes 6, 47-60, Nice 1995. Pannonien. - In: PAPP, A., JÄMBOR, Ä. & STEININGER, F.F. (Eds.): HRUSECKY, I., SEFARA, J., MASARYK, P. & LINTNEROVÄ, O.: The structu­ Chronostratigraphie und Neostratotypen. Miozän der Zentralen ral and facies development and exploration potential of the Slovak Paratethys VII, M6, Pannonien, 276-339, Akademiai Kiadö, part of the Danube Basin. - In: WESSELY, G. & LIEBL, W. (Eds.): Oil Budapest 1985. and gas in Alpidic thrustbelts and basins of Central and Eastern SAUERZOPF, F.: Beitrag zur Entwicklungsgeschichte des südburgen- Europe. EAGE (European Association of Geoscientists and ländischen Pannons. - Burgenländische Heimatblätter, 14, 1-16, Engineers) Special Publication 5, AM-A2B, Geological Society Eisenstadt 1952. Publishing House, Bath 1996. STOLIZCKA, F.: Kenntniss der Molluskenfauna der Cerithien- und JÄMBOR, Ä.: A Dunäntüli-közephegyseg pannöniai kepzödmenyei Inzersdorfer Schichten des ungarischen Tertiärbeckens. - (Pannonian in the Transdanubian Central Mountains). - Annals of Verhandlungen der k. k. zoologisch-botanischen Gesellschaft in the Hungarian Geological Institute, 62, 1-259, Budapest 1980. Wien, 7-10, Wien 1862. JIRICEK, R.: Die slowakische Donautiefebene. - In: PAPP, A., JÄMBOR, Ä. STRAUSZ, L: Das Pannon des mittleren Westungarns. - Annales & STEININGER, F.F. (Eds.): Chronostratigraphie und Neostratotypen. Historico-Naturales Musei Nationalis Hungarici, pars Mineralogica, Miozän der Zentralen Paratethys VII, M6, Pannonien, 276-339, Geologica et Palaeontologica, 5, 1-102, Budapest 1942. Akademiai Kiadö, Budapest 1985. SZILAJ, R., SZÖNOKY, M., MÜLLER, P., GEARY, D.H. & MAGYAR, I.: KEITH, J.F., VASS, D. & KovÄc, M.: The Danube lowland basin. - ESRI Stratigraphy, paleoecology, and paleogeography of the "Congeria Occasional Publication, New Series No. 11A-B, 63-87, Columbia ungulacaprae beds" (= Lymnocardium ponticum Zone) in NW 1994. Hungary: study of the Däka outcrop. - Acta Geologica Hungarica, KÖRÖSSY, L: A kisalföldi köolaj-es földgäzkutatäs földtani eredmenyei 42, 33-55, Budapest 1999. (Hydrocarbon geology of the Little Plain in Hungary). - Ältalänos TARI, G.: Alpine Tectonics of the Pannonian Basin. - Unpublished Földtani Szemle, 22, 99-174, Budapest 1987. Ph.D. thesis, Rice University, 501 p., Houston 1994. KORPÄS-HÖDI, M.: Palaeoecology and biostratigraphy of the TARI, G.: Neoalpine tectonics of the Danube Basin (NW Pannonian Pannonian Mollusca fauna in the northern foreland of the Basin, Hungary). - In: ZIEGLER, P.A. & HORVÄTH, F. (Eds.), Peri- Transdanubian Central Range. - Annals of the Hungarian Tethys Memoir 2: Structure and Prospects of Alpine Basins and Geological Institute, 66, 1-163, Budapest 1983. Forelands. Memoires du Museum national d'Histoire naturelle, 170, KORPÄS-HÖDI, M.: A Szombathely II. sz. füräs pannöniai (s. I.) mollu- 439-454 +enclosures 1-3, Paris 1996. scäi (The Pannonian (s. I.) molluscs of borehole section VITALIS, I.: A Limnocardium vaho-costatum n. sp. - Mathematischer Szombathely II). - Annual Report of the Hungarian Geological und Naturwissenschaftlicher Anzeiger der Ungarischen Akademie Institute of 1990, 505-525, Budapest 1992. der Wissenschaften, 51, 696-704, Budapest 1934. KovÄc, M. & BARÄTH, I.: Tektonicko-sedimentamy vyvoj alpsko-karpat- VITÄLIS, I.: Sopron környekenek szarmäciai es pannöniai-pontusi üle- sko-panönskej stycnej zöny pocas miocenu (Tectono-sedimentary dekei es kövületei (Les Sedimentes et fossils sarmatiens et panno- development of the Alpine-Carpathian-Pannonian junction zone du­ no-pontiens des environs de Sopron). - Annals of the Hungarian ring the Miocene). - Mineralia Slovaca, 28,1-11, Bratislava 1996. Geological Institute, 40, 3-75, Budapest 1951. LANTOS, M. & ELSTON, D.P.: Low- to high-amplitude oscillations and secular variation in a 1.2 km late Miocene inclination record. - Physics APPENDIX of the Earth and Planetary Interiors, 90, 37-53, Amsterdam 1995. LUEGER, J. P.: Die Molluskenfauna aus dem Pannon (Obermiozän) des Fölligberges (Eisenstädter Bucht) im Burgenland (Österreich). In the Appendix we give known occurrences of characteristic bi­ - Mitteilungen der österreichischen geologischen Gesellschaft, 73, valves of sandy, littoral deposits of Lake Pannon from Burgenland 95-134, Wien 1980. and the Transdanubian Central Range, respectively. Species names MAGYAR, I.: Mollusc fauna and flora of the Pannonian quartz sandstone given by the collector or author of paper are given in brackets where at Mindszentkälla, Hungary. - Annales Universitatis Scientiarum they do not correspond to our identification. Key to the collections Budapestinensis de Rolando Eotvos nominatae, sectio geologica, is given in the Materials and methods section. Materials from out­ 28, 209-222, Budapest 1988. crops in the vicinity of Müllendorf and Großhöflein (Föllig or Fölick) are united under the name "Großhöflein". MAGYAR, I., GEARY, D.H., SÜTÖ-SZENTAI, M., LANTOS, M. & MÜLLER, P.: Integrated biostratigraphic, magnetostratigraphic and chronostratig­ Because most of the MÄFI collection was obtained before 1920 by field raphie correlations of the Late Miocene Lake Pannon deposits. - geologists of the Royal Hungarian Geological Institute, this material is ori­ Acta Geologica Hungarica, 42, 5-31, Budapest 1999. ginally labelled and registered according to old Hungarian settlement names. In order to facilitate identification, we give these names in parenthe­ MARINESCU, Fl.: Le faciostratotype de Rädmäne§ti (Banat roumain). - sis after their Slovak and German counterparts at the end of the Appendix. In: STEVANOVIC, P.M., NEVESSKAJA, L.A., MARINESCU, Fl., SOKAC, A. & JÄMBOR, Ä. (Eds.): Chronostratigraphie und Neostratotypen. Neogen der Westlichen ("Zentrale") Paratethys VIII, PH, Pontien, The Burgenland fauna 436-439. JAZU and SANU, Zagreb-Beograd 1990. Unio atavus MATTICK, R.E., TELEKI, P.G., PHILLIPS, R.L., CLAYTON, J.L., DAVID, Gy., Großhöflein: LUEGER, 1980 [Psilunio atavus]; JPL; FS; MÄFI POGÄCSÄS, Gy., BARDÖCZ, B. & SIMON, E.: Structure, stratigraphy, Stegersbach: SAUERZOPF, 1952 [Psilunio stegersbachensis]; DHG and petroleum geology of the Little Plain basin, northwestern Olbendorf: SAUERZOPF, 1952 [Psilunio stegersbachensis]

676 Litzelsdorf: FS [Psilunio litzelsdorfensis] Lymnocardium subdiprosopum Oberdorf: SAUERZOPF, 1952 [Psilunio stegersbachensis]; FS; MÄFI Großhöflein: LUEGER, 1980 [L. diprosopum]; JPL; FS Pezinok: MÄFI "Lymnocardium" carnuntinum Congeria spathulata Großhöflein: DHG Großhöflein: LUEGER, 1980; JPL; FS : MÄFI [Limnocardium pseudosuessi] Stegersbach: DHG; FS : MÄFI Oberdorf: FS "Lymnocardium" danicici Congeria ungulacaprae Großhöflein: LUEGER, 1980; JPL; MÄFI; DHG; FS Pezinok: FORDINÄL, 1997; MM Sopron: TTM [Limnocardium galeatum] Sopran: ANDRUSOV, 1897 [Congeria hoemesi]; PAPP, 1953, 1985 [Congeria hoemesi]; MÄFI "Lymnocardium" stpjadinovici Unterschützen: MÄFI Congeria pancici Stegersbach: MÄFI Großhöflein: LUEGER, 1980; JPL; FS Stegersbach: SAUERZOPF, 1952 [Congeria pancici longiconcha]; PAPP, "Lymnocardium" stoosi, late form 1953, 1985; FS; DHG; MÄFI; BL St. Margarethen im Burgenland: MÄFI Steinbach: MÄFI Bratislava: FORDINÄL, 1995 [Pseudocatillus simplex] : MÄFI; FS Stegersbach: MÄFI, DHG St. Margarethen im Burgenland: MÄFI Großhöflein: LUEGER, 1980 [Pseudocatillus simplex]; DHG; FS Siegendorf: FS Congeria praebalatonica Burgau: FS Großhöflein: LUEGER, 1980 [Congeria balatonica]; JPL; FS Stegersbach: SAUERZOPF, 1952 [ Congeria spathulata praebalatonica]; Caladacna steindachneri PAPP, 1953 [Congeria spathulata praebalatonica] Großhöflein: JPL Litzelsdorf: SAUERZOPF, 1952 [Congeria spathulata praebalatonica] Sopron: TTM Bratislava: FORDINÄL, 1995 [Caladacna ornata biseptä] Congeria doderleini Stegersbach: PAPP, 1953, 1985; DHG Euxinicardium schreteri Pezinok: FORDINÄL, 1991, 1997 Stegersbach: FS; DHG; MÄFI St. Margarethen im Burgenland: FS "Didacna" deserta Congeria gitneri Siegendorf: MÄFI Großhöflein: LUEGER, 1980 [Dreissena minima]; DHG Großhöflein: LUEGER, 1980; JPL Stegersbach: DHG Stegersbach: STOLICZKA, 1862 [Cardium desertum]; PAPP, 1953, 1985; FS; DHG; MÄFI Dreissena bipartita Steinbach: MÄFI Pezinok: FORDINÄL, 1997 Sopron: TTM Siegendorf: JPL Lymnocardium schedelianum Brück an der Leitha: MÄFI Großhöflein: LUEGER, 1980; JPL; FS Bratislava: FORDINÄL, 1995 Bratislava: FORDINÄL, 1995 St. Margarethen im Burgenland: FS Stegersbach: DHG; FS St. Margarethen im Burgenland: MÄFI; FS "Monodacna" viennensis Zillingtal: MÄFI Großhöflein: JPL Oggau: FS Stegersbach: MÄFI

Lymnocardium brunnense Parvidacna petkovici Großhöflein: JPL Stegersbach: FS [Parvidacna loerentheyi]; PAPP, 1953,1985; NHMW Sopran: TTM Bratislava: FORDINÄL, 1993 St. Margarethen im Burgenland: MÄFI; FS Großhöflein: LUEGER, 1980 Bratislava: FORDINÄL, 1995

Lymnocardium conjungens Großhöflein: LUEGER, 1980; JPL; MÄFI The TCR fauna Stegersbach: FS; DHG; MÄFI; TTM Steinbach: MÄFI Unio mihanovici Sopran: TTM Däka: SZILAJ et al., 1999; MM Gols: JPL Päpa: MÄFI St. Margarethen im Burgenland: MÄFI; FS Kocs: MÄFI Pezinok: FORDINÄL, 1997; MÄFI; MM Romänd: UV Siegendorf: NHMW Ukk: MÄFI Fertöräkos: TTM Veszpremgalsa: MÄFI Bratislava: FORDINÄL, 1995 Kerekteleki: MÄFI Purbach am Neusiedlersee: FS Oggau: FS Congeria äff. simulans turgida Neszmely: ANDRUSOV, 1897 [Congeria turgida] Lymnocardium hantkeni Däka: SZILAJ et al., 1999; MM; MÄFI Stegersbach: NHMW; FS [Lymnocardium stegersbachensis n. sp.] Päpa: MÄFI Großhöflein: LUEGER, 1980; JPL Ukk: MÄFI

Lymnocardium edlaueri Congeria ungulacaprae Großhöflein: JPL Kup: FUCHS, 1870 [Congeria balatonicavar. crassitestä]; MÄFI Stegersbach: FS; DHG Mocsa: KORPÄS-HÖDI, 1983 Pezinok: MÄFI [Limnocardium vicinum]; MM Tata: KORPÄS-HÖDI, 1983 St. Margarethen im Burgenland: FS Chl'aba: MÄFI Gols: JPL Somlöväsärhely: ANDRUSOV, 1897; MÄFI Doba: MÄFI Lymnocardium tucani Gic: MÄFI Großhöflein: LUEGER, 1980; JPL; DHG; MÄFI; FS Kapolcs: MÄFI Stegersbach: FS; DHG; MÄFI Neszmely: MÄFI (?) Sopran: TTM Kocs: MÄFI Kömlöd: MÄFI Päpa: MÄFI Környe: MÄFI Somlöjenö: STRAUSZ, 1942 Kisber: MÄFI Tüskevär: MÄFI Csep: MÄFI Csäszär: MÄFI Lymnocardium ponticum Agostyän: MÄFI Däka: SZILAJ et al., 1999; MM Kerekteleki: MÄFI Nyäräd: MM Romänd: UV; MÄFI Naszäly: KORPÄS-HÖDI, 1983 [Limnocardium trifkovici, L decorum] Päpa: MÄFI Kocs: KORPÄS-HÖDI, 1983 Padragküt: MÄFI Päpa: MÄFI Käptalantöti: MÄFI Kup: NHMW [Cardium decorum]; BRUSINA, 1902 [Limnocar­ dium trifkovici]; HPM Dreissenomya unioides Läzi: BARTHA, 1963; MÄFI Lymnocardium tucani Däka: SZILAJ et al., 1999; MM; MÄFI Päpa: STRAUSZ, 1942 [Limnocardium banaticum var.] Szäk: MM Läzi: MÄFI [Limnocardium sp.] Tüskevär: MÄFI Dreissena auricularis Kup: FUCHS, 1870; MÄFI "Lymnocardium" priscae Däka: SZILAJ et al., 1999; MM; MÄFI Päpa: STRAUSZ, 1942; MAFI Päpa: MÄFI Läzi: MÄFI Läzi: BARTHA, 1963; MÄFI Tüskevär: STRAUSZ, 1942 Naszäly: KORPÄS-HÖDI, 1983 (?) Kocs: KORPÄS-HÖDI, 1983 Dunaszentmiklös: KORPÄS-HÖDI, 1983 Kup: HPM Szäk: MM Somlöjenö: UV "Lymnocardium" cf. wurmbi Veszpremgalsa: MÄFI Däka: SZILAJ et al., 1999; MM; MÄFI Päpa: TTM Dreissena bipartita Kup: ANDRUSOV, 1897; BRUSINA, 1902; HPM Caladacna steindachneri Däka: SZILAJ et al., 1999; MM Däka: STRAUSZ, 1942; SZILAJ et al., 1999; MM; TTM Szäk: MM Lymnocardium variocostatum Päpa: STRAUSZ, 1942; MÄFI Läzi: BARTHA, 1963, MÄFI Dunaalmäs: KORPÄS-HÖDI, 1983 Däka: SZILAJ et al., 1999; MM; MÄFI; TTM Tüskevär: MÄFI Kocs: VIT ALIS, 1934; MÄFI Kup: MÄFI Euxinicardium schreteri Päpa: MÄFI; UV Däka: SZILAJ et al., 1999; MM; MAFI Tärkäny: MÄFI Läzi: MÄFI Somlöjenö: UV; MÄFI Szäk: MM Romänd: UV Päpa: MÄFI Tüskevär: TTM Somlöjenö: STRAUSZ, 1942 Gic: MÄFI Tüskevär: MÄFI Külsövat: MÄFI Veszpremgalsa: MAFI "Didacna" deserta Kup: MILAN et al., 1974 [Didacna chyzeri]; HPM Lymnocardium apertum (?) Neszmely: MÄFI Läzi: MÄFI Däka: SZILAJ et al., 1999; MM; MÄFI; TTM "Monodacna" viennensis Szäk: MM Däka: MM Dunaalmäs: KORPÄS-HÖDI, 1983 Läzi: MÄFI Päpa: MÄFI Veszpremgalsa: MÄFI Parvidacna sp. Läzi: BARTHA, 1963; MÄFI Lymnocardium penslii Neszmely: KORPÄS-HÖDI, 1983 [Parvidacna planicostatä] Läzi: BARTHA, 1963; MÄFI Kocs: KORPÄS-HÖDI, 1983 [Parvidacna planicostatä] Däka: SZILAJ et al., 1999; MM; MÄFI; TTM; Dunaalmäs: KORPÄS-HÖDI, 1983 [Parvidacna planicostatä] Kocs: KORPÄS-HÖDI, 1983; MAFI Päpa: MÄFI Szäk: MM Däka: MM Päpa: MÄFI Kup: HPM Veszpremgalsa: MÄFI Ukk: MAFI Hungarian settlement names from Austria and Slovakia in the Somlöjenö: MÄFI; UV MÄFI collection (in parenthesis) Kup: NHMW; MÄFI Romänd: UV Brück an der Leitha (Bruk) Tüskever: TTM; MÄFI [Limnocardium banaticum] Chl'aba (Helemba) Neszmely: TTM; MÄFI Gols (Gälos) Dunaalmäs: TTM Großhöflein (Nagyhöfläny) Bärsonyos: MÄFI Litzelsdorf (Lödös) Tärkäny: MÄFI Oberdorf (Örälläs) Gic: MÄFI Olbendorf (Ober) Kapolcs: MÄFI Pezinok (Bazin) Kerekteleki: MÄFI St. Margarethen im Burgenland (Szentmargitbanya) Päpakoväcsi: MÄFI Siegendorf (Cinfalva) Stegersbach (Szentelek) Lymnocardium hantkeni Steinbach (Köpatak) Kup: FUCHS, 1870; MÄFI Steinbrunn (Büdösküt) Szäk: MM; MÄFI Unterschützen (Alsölövö) Däka: SZILAJ et al., 1999; MM Zillingtal (Völgyfalu)

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